Peer-reviewed publications

Reliability of TMS Phosphene Threshold Estimation: Toward a Standardized Protocol
Mazzi C., Savazzi, S., Abrahamyan, A., Ruzzoli, M. (2017)
Brain Stimulation. Available online 2 February. More Abstract.Background
Phosphenes induced by transcranial magnetic stimulation (TMS) are a subjectively described visual phenomenon employed in basic and clinical research as index of the excitability of retinotopically organized areas in the brain.
Objective
Phosphene threshold estimation is a preliminary step in many TMS experiments in visual cognition for setting the appropriate level of TMS doses; however, the lack of a direct comparison of the available methods for phosphene threshold estimation leaves unsolved the reliability of those methods in setting TMS doses. The present work aims at fulfilling this gap.
Methods
We compared the most common methods for phosphene threshold calculation, namely the Method of Constant Stimuli (MOCS), the Modified Binary Search (MOBS) and the Rapid Estimation of Phosphene Threshold (REPT). In two experiments we tested the reliability of PT estimation under each of the three methods, considering the day of administration, the participants' expertise in phosphene perception and the sensitivity of each method to the initial values used for the threshold calculation.
Results
We found that MOCS and REPT have comparable reliability when estimating phosphene thresholds, while MOBS estimations appear less stable.
Conclusions
Based on our results, researchers and clinicians can estimate phosphene threshold according to MOCS or REPT equally reliably, depending on their specific investigation goals. We suggest several important factors for consideration when calculating phosphene thresholds and describe strategies to adopt in experimental procedures.
http://dx.doi.org/10.1016/j.brs.2017.01.582PDF

Adaptable History Biases in Human Perceptual Decisions
Abrahamyan, A., Silva L. L., Dakin S. C., Carandini M., & Gardner J. L. (2016)
Proceedings of the National Academy of Sciences, 113(25), E3548–57. More Abstract.
When making choices under conditions of perceptual uncertainty, past experience can play a vital role. However, it can also lead to biases that worsen decisions. Consistent with previous observations, we found that human choices are influenced by the success or failure of past choices even in a standard two-alternative detection task, where choice history is irrelevant. The typical bias was one that made the subject switch choices after a failure. These choice-history biases led to poorer performance and were similar for observers in different countries. They were well captured by a simple logistic regression model that had been previously applied to describe psychophysical performance in mice. Such irrational biases seem at odds with the principles of reinforcement learning, which would predict exquisite adaptability to choice history. We therefore asked whether subjects could adapt their irrational biases following changes in trial order statistics. Adaptability was strong in the direction that confirmed a subject’s default biases, but weaker in the opposite direction, so that existing biases could not be eradicated. We conclude that humans can adapt choice history biases, but cannot easily overcome existing biases even if irrational in the current context: adaptation is more sensitive to confirmatory than contradictory statistics.
doi.org/10.1073/pnas.1518786113PDF

Mimicry and Expressiveness of an ECA in Human-Agent Interaction: Familiarity Breeds Content!
Stevens, C. J., Pinchbeck B., Lewis, T., Luerssen, M., Pfitzner, D., Powers, D., Abrahamyan, A., Leung, Y., & Gibert, G. (2016)
Computational Cognitive Science, 2(1), 1-14. More Abstract.Background.
Two experiments investigated the effect of features of human behaviour on the quality of interaction with an Embodied Conversational Agent (ECA).
Methods.
In Experiment 1, visual prominence cues (head nod, eyebrow raise) of the ECA were manipulated to explore the hypothesis that likeability of an ECA increases as a function of interpersonal mimicry. In the context of an error detection task, the ECA either mimicked or did not mimic a head nod or brow raise that humans produced to give emphasis to a word when correcting the ECA’s vocabulary. In Experiment 2, presence versus absence of facial expressions on comprehension accuracy of two computer-driven ECA monologues was investigated.
Results.
In Experiment 1, evidence for a positive relationship between ECA mimicry and lifelikeness was obtained. However, a mimicking agent did not elicit more human gestures. In Experiment 2, expressiveness was associated with greater comprehension and higher ratings of humour and engagement.
Conclusion.
Influences from mimicry can be explained by visual and motor simulation, and bidirectional links between similarity and liking. Cue redundancy and minimizing cognitive load are potential explanations for expressiveness aiding comprehension.
doi.org/10.1186/s40469-016-0008-2

Low Intensity TMS Enhances Perception of Visual Stimuli
Abrahamyan, A., Clifford, C. W. G., Arabzadeh, E., & Harris, J. A. (2015)
Brain Stimulation, 8(6), 1175–1182. More Abstract.Background
Transcranial magnetic stimulation (TMS) is a popular functional mapping tool in cognitive and sensory neuroscience. While strong TMS typically degrades performance, two recent studies have demonstrated that weak TMS, delivered to visual cortex, can improve performance on simple visual tasks. The improvement was interpreted as the summation of visually-evoked and TMS-elicited neuronal activity in visual cortex, but the nature of this interaction remains unclear.
Objective
The present experiments sought to determine whether these weak pulses of TMS assist subjects to see the visual stimulus itself or create a distinct “melded” percept that may not be recognizable as the visual stimulus.
Methods
We measured contrast thresholds in an orientation discrimination task in which participants reported the orientation (left or right) of gratings tilted 45° from vertical.
Results
Weak TMS improved sensitivity for identifying gratings, suggesting that TMS sums with but preserves orientation information so that the subject can recognize the visual stimulus. We explain the effect using a mechanism of non-linear transduction of sensory signals in the brain.
Conclusions
The capability of low-intensity TMS to augment the neural signal while preserving information encoded in the stimulus can be employed as a novel approach to study the neural correlates of consciousness by selectively “pushing” an unconscious stimulus into consciousness.
http://doi.org/10.1016/j.brs.2015.06.012PDF

Brain-stimulation Induced Blindsight: Unconscious Vision or Response Bias?
Lloyd, D. A., Abrahamyan, A., & Harris, J. A. (2013)
PLoS ONE, 8(12), e82828. More Abstract. A dissociation between visual awareness and visual discrimination is referred to as “blindsight”. Blindsight results from loss of function of the primary visual cortex (V1) which can occur due to cerebrovascular accidents (i.e. stroke-related lesions). There are also numerous reports of similar, though reversible, effects on vision induced by transcranial Magnetic Stimulation (TMS) to early visual cortex. These effects point to V1 as the “gate” of visual awareness and have strong implications for understanding the neurological underpinnings of consciousness. It has been argued that evidence for the dissociation between awareness of, and responses to, visual stimuli can be a measurement artifact of the use of a high response criterion under yes-no measures of visual awareness when compared with the criterion free forced-choice responses. This difference between yes-no and forced-choice measures suggests that evidence for a dissociation may actually be normal near-threshold conscious vision. Here we describe three experiments that tested visual performance in normal subjects when their visual awareness was suppressed by applying TMS to the occipital pole. The nature of subjects’ performance whilst undergoing occipital TMS was then verified by use of a psychophysical measure (d') that is independent of response criteria. This showed that there was no genuine dissociation in visual sensitivity measured by yes-no and forced-choice responses. These results highlight that evidence for visual sensitivity in the absence of awareness must be analysed using a bias-free psychophysical measure, such as d', In order to confirm whether or not visual performance is truly unconscious.
http://doi.org/10.1371/journal.pone.0082828PDF

Improving Visual Sensitivity with Subthreshold Transcranial Magnetic Stimulation
Abrahamyan, A., Clifford, C. W. G., Arabzadeh, E., & Harris, J. A. (2011)
Journal of Neuroscience, 31(9), 3290–3294 More Abstract. We probed for improvement of visual sensitivity in human participants using transcranial magnetic stimulation (TMS). Stimulation of visual cortex can induce an illusory visual percept known as a phosphene. It is known that TMS, delivered at intensities above the threshold to induce phosphenes, impairs the detection of visual stimuli. We investigated how the detection of a simple visual stimulus is affected by TMS applied to visual cortex at or below the phosphene threshold. Participants performed the detection task while the contrast of the visual stimulus was varied from trial to trial according to an adaptive staircase procedure. Detection of the stimulus was enhanced when a single pulse of TMS was delivered to the contralateral visual cortex 100 or 120 ms after stimulus onset at intensities just below the phosphene threshold. No improvement in visual sensitivity was observed when TMS was applied to the visual cortex in the opposite hemisphere (ipsilateral to the visual stimulus). We conclude that TMS-induced neuronal activity can sum with stimulus-evoked activity to augment visual perception.
http://doi.org/10.1523/JNEUROSCI.6256-10.2011

The effect of TMS on Visual Motion Sensitivity: An Increase in Neural Noise or a Decrease in Signal Strength?
Ruzzoli, M., Abrahamyan, A., Clifford, C. W. G., Marzi, C. A., Miniussi, C., & Harris, J. A. (2011)
Journal of Neurophysiology, 106(1), 138–143. More Abstract The underlying mechanisms of action of transcranial magnetic stimulation (TMS) are still a matter of debate. TMS may impair a subject's performance by increasing neural noise, suppressing the neural signal, or both. Here, we delivered a single pulse of TMS (spTMS) to V5/MT during a motion direction discrimination task while concurrently manipulating the level of noise in the motion stimulus. Our results indicate that spTMS essentially acts by suppressing the strength of the relevant visual signal. We suggest that TMS may induce a pattern of neural activity that complements the ongoing activation elicited by the sensory signal in a manner that partially impoverishes that signal.
http://doi.org/10.1152/jn.00746.2010

Accurate and Rapid Estimation of Phosphene Thresholds (REPT)
Abrahamyan, A., Clifford, C. W. G., Ruzzoli, M., Arabzadeh, E., & Harris, J. A. (2011)
PLoS ONE, 6(7) e22342 More Abstract. To calibrate the intensity of transcranial magnetic stimulation (TMS) at the occipital pole, the phosphene threshold is used as a measure of cortical excitability. The phosphene threshold (PT) refers to the intensity of magnetic stimulation that induces illusory flashes of light (phosphenes) on a proportion of trials. The existing PT estimation procedures lack the accuracy and mathematical rigour of modern threshold estimation methods. We present an improved and automatic procedure for estimating the PT which is based on the well-established Ψ Bayesian adaptive staircase approach. To validate the new procedure, we compared it with another commonly used procedure for estimating the PT. We found that our procedure is more accurate, reliable, and rapid when compared with an existing PT measurement procedure. The new procedure is implemented in Matlab and works automatically with the Magstim Rapid2 stimulator using a convenient graphical user interface. The Matlab program is freely available for download.
PDFhttp://doi.org/10.1371/journal.pone.0022342